﻿A revision of the genus Ecpyrrhorrhoe Hübner, 1825 from China based on morphology and molecular data, with descriptions of five new species (Lepidoptera, Crambidae, Pyraustinae)

﻿Abstract The genus Ecpyrrhorrhoe Hübner, 1825 is revised. Most type materials were examined, and a preliminary phylogeny is presented based on a combined dataset of COI, 16S rRNA, 28S rRNA and EF-1α gene regions. The tree topology and morphological characters suggest that Paliga Moore, 1886 is a new synonym of Ecpyrrhorrhoe, and Yezobotys Munroe & Mutuura, 1969 is restored as a valid genus. According to the morphological evidence and tree topology, 17 species are recorded, including five new species described from China: E.allochroa Zhang & Xiang, sp. nov., E.rosisquama Xiang & Zhang, sp. nov., E.exigistria Zhang & Xiang, sp. nov., E.brevis Zhang & Xiang, sp. nov. and E.longispinalis Zhang & Xiang, sp. nov. Seven new combinations are created, E.damastesalis (Walker, 1859), comb. nov., E.minnehaha (Pryer, 1877), comb. nov., E.obliquata (Moore, 1888), comb. nov., E.rufipicta (Butler, 1880), comb. nov., E.fimbriata (Moore, 1886), comb. nov., E.machoeralis (Walker, 1859), comb. nov., and E.rubellalis (Snellen, 1890), comb. nov., as well as eight new synonyms, namely Leucocraspedaauratalis Warren, 1895, syn. nov., Pioneaschenklingi Strand, 1918, syn. nov., Paligarubicundalis Warren, 1896, syn. nov., E.angustivalvaris Gao, Zhang & Wang, 2013, syn. nov., Pyraustapygmaealis South, 1901, syn. nov., E.multispinalis Gao, Zhang &Wang, 2013, syn. nov., E.aduncis Gao, Zhang & Wang, 2013, syn. nov., and E.ruidispinalis Zhang, Li & Wang, 2004, syn. nov. All adults and their genital structures are illustrated and an identification key based on adult external morphology and genitalia is provided.

There are striking apomorphic characters available to diagnose species of Ecpyrrhorrhoe. These are a narrowly lanceolate uncus, long dorsolateral arms of the juxta, the presence of spines on the anellus, a slender longitudinal sclerite located in the posterior part of the ductus bursae, and a second (posterior) signum with spines in the female genitalia. Paliga Moore, 1886 shares some of these characters with species of Ecpyrrhorrhoe, but Yezobotys does not share these characters. Therefore, the relationship of Yezobotys and Paliga with Ecpyrrhorrhoe needs to be resolved.
Upon examination of pyraustine collections from China, and type specimens and other material from the Natural History Museum, London, United Kingdom, and the Senckenberg Entomological Institute, Brandenburg, Germany, some known species, and undescribed species were found to agree with the circumscription of Ecpyrrhorrhoe based on genitalia characters. In order to evaluate the generic placements of these species and the taxonomic composition of Ecpyrrhorrhoe, the phylogenetic relationships of Ecpyrrhorrhoe were studied with molecular data.

Molecular phylogenetic analysis
In total 24 species were included in the molecular phylogenetic analysis (Table 1), including the type species of Yezobotys and Paliga, five new species and six putative new combinations. Euclasta stoetzneri (Caradja, 1927) was chosen as the outgroup because it has been inferred as sister-group of the Pyraustini and Portentomorphini in Pyraustinae (Mally et al. 2019). Two species of Pyrausta Schrank, 1802, two species of Pagyda Walker, 1859, and two species of Anamalaia Munroe & Mutuura, 1969 were included as related taxa because of the similar external and genital characters, and a previous taxonomic treatment of Ecpyrrhorrhoe as a subgenus of Pyrausta (Hannemann 1964).
Total DNA was extracted from two legs and sometimes additionally from the abdomen of the dry specimens using the TIANGEN DNA extraction kit following the manufacturer's instructions. The nucleotide sequences of two mitochondrial genes, cytochrome c oxidase subunit I (COI) and 16S ribosomal RNA (16S rRNA), and two nuclear genes, 28S ribosomal RNA (28S rRNA) and Elongation factor-1 alpha (EF-1α) were selected for study. Primers used in this study and all PCRs performed follow Zhang et al. (2020). PCR products were confirmed with 1.5% agarose gel electrophoresis in Table 1. Species sampled for the molecular phylogenetic analyses; all species sequenced in this study except Euclasta stoetzneri, which was sequenced by Zhang et al. (2020). TAE buffer, then were direct-sequenced at Majorbio Bio-pharm Technology Co., Ltd (Guangzhou), utilizing the same primers used for PCR amplification. The sequences were aligned using Clustal W (Thompson et al. 1994) in MEGA 6 (Tamura et al. 2013) with default settings. The aligned matrix was corrected by eye. Gaps were treated as missing data. Phylogenetic analyses were inferred using Bayesian inference (BI) method in MrBayes 3.2.6 (Ronquist et al. 2012) and maximum likelihood (ML) in RAxML 8.2.10 (Stamatakis 2014). BI analysis was run with independent parameters all under the GTR + G + I model for four gene partitions, as suggested by jModelTest 0.1.1 (Posada 2008). Two independent runs, each with four Markov Chain Monte Carlo (MCMC) simulations, were performed for 20 million generations sampled every 1000 th generation. The first 25% trees were discarded as burn-in, and posterior probabilities (PP) were determined from remaining trees. The ML analysis was executed under the GTR + G + I model for all gene partitions and with 1000 iterations for the bootstrap test. The bootstrap value (BS) ≥ 90 is considered absolute support, 75 ≤ BS < 90 is considered strong support, and 50 ≤ BS < 75 is considered weak support. PP ≥ 95 is considered strong support and 80 ≤ PP < 95 is considered weak support. The pairwise Kimura 2-Parameter (K2P) distances between species were calculated from the COI gene using MEGA 6 (Tamura et al. 2013).

Morphological analysis
The specimens studied, including the types of the newly described species, are deposited in the Museum of Biology, Sun Yat-sen University, Guangzhou, China (SYSBM), except for those held at the following institutions: Insect Collection of the College of Life Sciences, Nankai University, China (NKU), Natural History Museum, London, United Kingdom (NHMUK) and Senckenberg Deutsches Entomologisches Institut, Brandenburg, Germany (SDEI). Slides of dissected genitalia were prepared according to the protocols of Robinson (1976) and Li and Zheng (1996). Terminology of genitalia follows Maes (1995), except for "phallus" and "colliculum" for which we follow Kristensen (2003). Images of the specimens were taken using a Canon EOS 80D camera provided with a Canon 100 mm macro lens. The genitalia photographs were taken using a Zeiss Axio Scope.A1 in combination with a Zeiss AxioCam camera and the Axio Vision SE64 program on a Windows PC. Source images were then aligned and stacked with Helicon Focus to obtain a composite image. All the pictures were edited using Adobe Photoshop SC5.
Both BI and ML analyses of the concatenated dataset inferred fully congruent relationships with only subtle differences in posterior probability and bootstrap values (Fig. 1). The tree topology indicates that Yezobotys Munroe & Mutuura, 1969 should be restored as a valid genus because the type species, Y. dissimilis (Yamanaka, 1958), appears as sister to Anamalaia (PP = 1, BS = 100). The monophyly of Ecpyrrhorrhoe is supported (PP = 0.99, BS = 63). The genus Pagyda Walker, 1859 is the sister group of Ecpyrrhorrhoe (PP = 0.99, BS = 67). Among Ecpyrrhorrhoe species included in the analysis, the majority of the basal nodes are strongly supported in BI but relatively poorly supported in ML.
According to the tree topology and morphological characters, the genus Ecpyrrhorrhoe can be divided into three species groups (The A clade, B clade, and C clade). The A clade is the sister group to B clade + C clade (PP = 0.99, BS = 63). The A clade is composed of E. allochroa and E. damastesalis. The B clade consists  The results of the molecular phylogenetic analyses support the placement of five undescribed species (named as E. allochroa sp. nov., E. rosisquama sp. nov., E. exigistria sp. nov., E. brevis sp. nov., and E. longispinalis sp. nov.) in Ecpyrrhorrhoe, the transfer of E. rubellalis (Snellen, 1890), comb. nov. from Pyrausta Schrank, 1802 to Ecpyrrhorrhoe, the transfer of E. obliquata (Moore, 1888), comb. nov. and E. fimbriata (Moore, 1886), comb. nov. from Anania Hübner, 1823 to Ecpyrrhorrhoe, and the transfer of E. damastesalis (Walker, 1859), comb. nov., E. rufipicta (Butler, 1880), comb. nov., andE. minnehaha (Pryer, 1877), comb. nov. from Paliga Moore, 1886 to Ecpyrrhorrhoe. The taxonomic details are given below. Diagnosis. The wings of species of Ecpyrrhorrhoe are usually yellow, sometimes decorated with pink or brown scales. In appearance, they are similar to some species of Pyrausta Schrank, 1802 and Pseudopagyda Slamaka, 2013, but can be distinguished by the usually obvious dark brown subterminal band on the underside of wings. They are characterized by the lanceolate, densely setose uncus; the mostly thumb-shaped sella extending to the ventral margin of the valva; the long arms of the bifid juxta, usually; usually the presence of several spines or sclerites on the anellus in the male genitalia. In the female genitalia, the strongly sclerotized antrum, the slender sclerite located in the posterior part of the ductus bursae, and the second (posterior) signum bearing spines are characteristic. Description. Frons oblique, smoothly scaled. Vertex with moderately raised scales projecting between antennae. Labial palpus porrect, second segment pointing obliquely upward, third segment pointing slightly downward; exceeding frons by approximately as much as length of head. Maxillary palpus small. Forewing termen gently arched. Hindwing frenulum single in male, with two acanthae in female. Wings usually yellow or yellowish brown, sometimes pink or covered with brown scales; forewing with antemedial and postmedial lines, orbicular and reniform stigma; underside of wings usually with obvious blackish brown subterminal band. Wing venation as in Fig. 2. Male genitalia. Uncus with lanceolate head and a nearly triangular, broad, naked base; densely covered with short simple, thick, setae, and every cluster of setae is made up of two independent setae. Transtilla inferior thin and stick-shaped. Valva elongated tongue-shaped; costa straight to concave; sella thumb-shaped or semicircular, extending to ventral margin of valva, usually with dense setae or spines; sacculus broad, usually with a wide dorsal protrusion. Dorsolateral arms of juxta usually long and tapering; anellus with several spines. Saccus nearly triangular. Phallus tubular.
Distribution. Asia, Europe, South Africa, Australia, North America.
Male genitalia (Fig. 22). Uncus with basal 2/5 nearly triangular and naked. Valva slightly curved, of even width, apex obtusely rounded; sella thumb-shaped and excurved, setose, bearing several spines on ventral margin; sacculus with dorsal 3/5 inflated into a nearly triangular protrusion. Juxta shield-shaped, with base wide, distal 1/4 bifid into thick and short arms with membranous extension; anellus with a long and curved, densely spinulose sclerite (attached to distal end of phallus in Fig. 22). Saccus rounded triangular. Phallus long and slender, distal part slightly curved upward and sclerotized, with a cluster of interlaced spicules on vesica; distal end with a bent, spine-like cornutus. Diagnosis. In the male genitalia ( Fig. 23), E. damastesalis can be characterized by the relatively thick and tapered uncus, the slender and straight sella bearing more strongly sclerotized short spines almost vertically placed on ventral and distal margins,   (Guangxi) 11 E. rubiginalis, female (Xinjiang) 12 E. machoeralis, type, female (Sri Lanka). Scale bars: 5.0 mm. and the three groups of spines present on anellus. The female genitalia (Fig. 38) are unique, readily separable by the triangular antrum, the wrinkled corpus bursae, the significantly large and generally subtriangular rhomboid signum with anterior and posterior parts asymmetrical and bearing a slightly curved carina, as well as the longer spines on markedly large second (posterior) signum.
Redescription (Figs 4,5). Head. Frons yellow, or yellowish brown scattered with rosy scales, with white lateral bands. Vertex pale yellow, usually scattered with rosy scales. Labial palpus yellowish brown or brown, usually scattered with rosy scales, contrastingly white at base ventrally. Maxillary palpus yellowish brown or brown, usually scattered with rosy scales, pale yellow terminally. Thorax. Dorsal side yellow, and ventral side white; tegula yellow or mixed with rosy scales sometimes. Wings. Forewing length: 10.0-14.0 mm. Forewing narrow and elongated; pale yellow, usually covered with rosy scales on veins, markings yellowish brown or rosy; antemedial line obliquely from 1/4 of costa to beyond posterior margin of cell, then deeply dentated to basal 1/3 of dorsum; orbicular stigma oval and distinct; reniform stigma comma-shaped, short and thick, sometimes concave; postmedial line bent inwards from 3/4 of costa, then arched and crenulated to basal half of CuA 1 , finally sharply dentated to 2/3 of posterior margin; subterminal line crenulated, sometimes faint; vein ends sometimes bearing triangular spots; fringe pale yellow, mostly mixed with rosy scales sometimes. Hindwing yellowish white, termen yellowish brown or rosy; fringe pale yellow, sometimes with some pink toward apex. Abdomen. Pale yellow dorsally, white ventrally.
Male genitalia (Fig. 23). Uncus relatively thick and tapered, with basal 1/3 nearly triangular and naked. Valva slightly broad, with costa nearly straight or concave and ventral margin curved, basal part weakly narrowed, remainder of even width, apex obtusely rounded; sella thumb-shaped and straight, extending inward, bearing short spines vertically settled on ventral and distal margins; sacculus with dorsal 3/5 inflated into a nearly triangular protrusion. Juxta shield-shaped, medially weakly sclerotized, bifid; anellus with three groups of spines (attached to distal juxta in Fig. 23). Saccus narrowly triangular. Phallus long and slender, basal part slightly curved, with a bunch of interlaced spicules on vesica.
Material examined. Type material. Type: 1♀ Remarks. The larvae of Ecpyrrhorrhoe damastesalis are leaf skeletonizers of teak (Tectona grandis). Severe infestations, causing 90%-100% defoliation, has been recorded from Malaysia and Guangdong, China (Intachat 1998;Lin et al. 2018). The misidentification of E. damastesalis as Ecpyrrhorrhoe machoeralis (Walker, 1859), comb. nov. is common and has been verified in Java and Thailand (Intachat 1998) and Hainan, China (Wu et al. 1977;Wang 1980 Diagnosis. This species can be differentiated from other species of the genus by its smaller forewing length (8.0-10.0 mm) and the yellow or rosy-red forewing usually bearing strongly contrasting spots and relatively smooth and slender lines (Figs 6, 7). In the male genitalia (Fig. 24), E. minnehaha is somewhat similar to E. fimbriata, but can be distinguished by the much more rounded ventral margin of valva, excurved sella densely bearing thick setae, pointed arms of juxta without tooth, and presence of a group of spines on the anellus. The female genitalia (Fig. 39) can be distinguished from congeners by the short cup-shaped antrum densely covered with minute spines.  (Figs 6, 7). Head. Frons yellow or yellowish brown with white lateral bands. Vertex pale yellow. Labial palpus yellow or yellowish brown, contrastingly white at base ventrally. Maxillary palpus yellow or yellowish brown, pale yellow terminally.

Redescription
Thorax. Dorsal side yellow or yellowish brown, ventral side white. Wings. Forewing length: 8.0-10.0 mm. Forewing yellowish brown, or rosy-red with posterior area straw yellow, markings blackish brown; antemedial line slightly arched from 1/4 of costa to 1/3 of posterior margin; orbicular stigma very small, dot-shaped, sometimes indistinct in rosy-red individuals; reniform stigma comma-shaped, nearly straight or weakly concave; postmedial line from 3/4 of costa to middle of CuA 1, bending to 1/3 of CuA 2 , then weakly curved to 2/3 of posterior margin; without distinct fuzzy patch posterolateral of cell indistinct; terminal line brown to dark brown; fringe with basal half brown to dark brown, distal half silver-white. Hindwing yellowish brown, with basal and subterminal area scattered with blackish brown scales, posterior area pale yellow; postmedial line from 2/3 of costa to 1/2 of CuA 1 , bending inward to 1/3 of CuA 2 , then weakly convex to 2/3 of posterior margin; the fuzzy patch posterolateral of cell blackish brown; terminal line and fringe as in forewing. Abdomen. Pale yellow dorsally, white ventrally.
Male genitalia (Fig. 24). Uncus with basal 1/3 nearly triangular and naked. Valva with costa slightly concave and ventral margin curved, with apex rounded; sella excurved, thumb-shaped, densely bearing thick setae; sacculus with distal 3/5 inflated into a broad and nearly triangular protrusion. Juxta with base wide, distal half bifid into slender, straight, and widely separated arms; anellus with a cluster of short spines (attached to distal end of phallus in Fig. 24). Saccus rounded triangular. Phallus long and moderately thick; cornuti presented as a bunch of spines, with a large spine distally and connected with a lanceolate sclerite at base.
Female genitalia (Fig. 39). Anterior apophysis ~ 2× length of posterior apophysis. Antrum cup-shaped, short and broad, strongly sclerotized, inner wall densely covered with minute spines; colliculum long, with two longitudinal ridges; ductus seminalis inserting to anterior end of colliculum and with a wide sclerite located opposite to colliculum; ductus bursae ~ 2× as long as length of corpus bursae, basal 2/3 bearing a slim sclerite. Corpus bursae oval; rhomboid signum narrow, with maximal length equal to half length of corpus bursae; second (posterior) signum small and somewhat curved, bearing short spines.
Material examined. Type material. Remarks. The type material of Pyrausta minnehaha Pryer, 1877 has rosy-red forewings bearing strongly contrasting spots and relatively smooth and slender lines, and the type series of Pionea schenklingi Strand, 1918 has the same appearance. Leucocraspeda auratalis Warren, 1895 has a yellowish brown forewing, but bears the same markings, hindwings and fringe as Pyrausta minnehaha and Pionea schenklingi. The genitalic characters indicate that these all belong to the same species. Although there is striking variation in this species, it is most easily identified by its appearance and its small body size. On the basis of currently available material, specimens with yellowish brown forewings are the commonest form in its range. (Moore, 1888) Diagnosis. In appearance, Ecpyrrhorrhoe obliquata is similar to E. exigistria in the brown subterminal area of the wings, as well as in the oblique, dark brown streak on forewing which distinguish them from all other species of the genus. Genitalia characters readily distinguish E. obliquata from other members of the genus in the semi-circular sella with basal half of ventral margin bearing curved spines. It can be best distinguished from E. exigistria by the larger forewing length (forewing length: 10.0-12.0 mm), the more distinct and longer streak of the forewing, in the male genitalia (Fig. 25) by the shape of sella, and the weakly sclerotized arms of the bifid juxta, which is without teeth, and in the female genitalia (Fig. 40) by the shape of antrum.

Ecpyrrhorrhoe obliquata
Redescription (Fig. 8). Head. Frons and vertex yellow, frons with lateral white bands. Labial palpus yellowish brown, contrastingly white at base ventrally. Maxillary palpus yellowish brown, white terminally. Thorax. Dorsal side yellow and ventral side white; tegula yellow with brown base. Wings. Forewing length: 10.0-12.0 mm. Forewing pale yellow, basal half of costal band and posterior half of subterminal area brown; stigmata and lines dark brown; antemedial line strongly oblique from 1/5 of costa to 1/2 of 1A, then dentated to 2/5 of posterior margin; orbicular stigma dotshaped, sometimes indistinct; reniform stigma comma-shaped, thick and straight; an oblique, dark brown streak from posterior end of reniform stigma weakly curved and extended to tornus; postmedial line obliquely outwards from 3/5 of costa to 1/2 of M 1 , then dentated inwards to 2/3 of posterior margin; terminal line dark brown and intermittent; fringe with basal half yellow and blackish brown, distal half with anterior half yellow and posterior half dark brown. Hindwing pale yellow, subterminal area from dark brown to brown, gradually paler to tornus; postmedial line dark brown, slightly dentate from 2/3 of M 1 to 1/2 of CuA 1 , then bending inwards to base of CuA 1 , finally undulated to 2/3 of posterior margin; terminal line and fringe as in forewing. Abdomen. Pale yellow dorsally, each segment with posterior margin whitish, black on distal end; white ventrally.
Male genitalia (Fig. 25). Uncus with basal 1/3 nearly triangular and naked. Valva with costa nearly straight or concave and ventral margin curved, basal part weakly narrowed, remainder of even width, apex obtusely rounded; sella nearly semi-circular, bearing short and curved spines on distal half of ventral margins; sacculus with middle part inflated into a thumb-shaped protrusion. Juxta with base wide, distal 3/4 bifid into pointed arms. Saccus narrowly triangular. Phallus long and straight, with a bundle of short spines assembling into cone-shape at distal end.
Female genitalia (Fig. 40). Anterior apophysis ~ 2× length of posterior apophysis. Antrum cup-shaped, with anterior 1/3 strongly sclerotized and covered with spinules on inner wall, posterior 1/3 abruptly broad and partly wrinkled; colliculum long; ductus seminalis connecting to anterior end of colliculum and with a short sclerite located opposite to colliculum; ductus bursae ~ 2.5-3× as long as diameter of corpus bursae, basal 1/3 bearing a slim sclerite. Corpus bursae globular; rhomboid signum with maximal length approximately 1/2 as long as diameter of corpus bursae; the second (posterior) signum nearly thumb-shaped bearing a wide base, sparsely covered with long spines.
Material examined. Type material.  (Moore, 1886) in having nearly the same forewing length, yellow wings bearing brown markings and an almost indistinct, brown subterminal band (Fig. 9). However, it can be differentiated from E. fimbriata by more dentated and relatively thick postmedial line on both wings, in the male genitalia (Fig. 26) by the nearly oval valva, the thumb-shaped dorsal sella, the triangular ventral sella bearing several spines, the longer and strongly sclerotized arms of the juxta, and two long and pointed spines located on anellus.
Male genitalia (Fig. 26). Uncus with basal 1/2 nearly triangular and naked. Valva with costa slightly convex and ventral margin curved, with basal part narrowed, remainder nearly oval, apex rounded; sella nearly triangular, bearing short spines on distal and inner margins, with a curved, finger-shaped dorsal projection; sacculus with middle part inflated into a triangular and setose protrusion. Juxta with base wide, distal 4/5 forming slender, long, and sclerotized arms dentate apically; anellus with two long and pointed spines (connected with distal arms of juxta in Fig. 26). Saccus rounded triangular. Phallus long and straight, cornuti presented as a narrow sclerite with dense and short spines.
Female genitalia (Fig. 41). Anterior apophysis ~ 2.5× length of posterior apophysis. Antrum cylindrical, tuberculate laterally on anterior end, strongly sclerotized and covered with spinules on inner wall; colliculum long and broad, narrower medially; ductus seminalis connecting to anterior end of colliculum and with a wide sclerite located opposite to colliculum; ductus bursae ~ 2.5-3× as long as diameter of corpus bursae, basal 2/5 bearing a slim sclerite. Corpus bursae globular; rhomboid signum with maximal length > 1/2 of diameter of corpus bursae; second (posterior) signum composed of a pair of round sclerites bearing dense and long spines. Remarks. Based on the substantial morphological similarity in the male genitalia between the types of Asopia rufipicta Butler, 1880 and Paliga rubicundalis Warren, 1896, Paliga rubicundalis is considered as a junior synonym of E. rufipicta (Butler). Diagnosis. Forewing length: 9.0-11.0 mm. Ecpyrrhorrhoe fimbriata is similar to E. rubiginalis both in appearance and in the male genitalia, but it can be differentiated from it by the relatively smooth and slender postmedial line on both wings (Fig. 10), in the male genitalia (Fig. 27), by the even width of the valva, dorsal projection of sella absent and the setose, thumb-shaped sella, the weakly sclerotized arms of the juxta with several short spines at apex, and the cluster of spines on the anellus (attached to distal end of phallus in Fig. 27 Gao, Zhang & Wang, 2013: 312. Syn. nov. Diagnosis. Forewing length: 9.0-12.0 mm. In appearance, Ecpyrrhorrhoe rubiginalis is similar to E. fimbriata, but the coloration of the wings and markings of E. rubiginalis is darker, and the patch in the hindwing is larger (Fig. 11). In the male genitalia (Fig. 28), the phallus of E. rubiginalis has a cluster of interlaced spicules and an oval sclerite bearing spines on the vesica, with three separate spines on the anellus (attached to distal end of phallus in Fig. 28 Wang & Sung, 1980: 305. Diagnosis. Forewing length: 8.0-10.0 mm. Ecpyrrhorrhoe machoeralis can be differentiated from congeners by the antrum with anterior half narrower than posterior half. Female genitalia (Fig. 43). Anterior apophysis ~ 1.5× length of posterior apophysis. Antrum long and tubular, with anterior half broad and slightly wrinkled, posterior half slightly narrow and as wide as colliculum; colliculum moderately long, somewhat difficult to differentiate from antrum; ductus seminalis connecting to anterior end of colliculum and with a weak sclerite located opposite to colliculum; ductus bursae ~ 2-2.5× as long as diameter of corpus bursae, basal 2/5 bearing a slim sclerite. Corpus bursae globular; rhombic signum with maximal length < ~ 1/2 length of diameter of corpus bursae; second (posterior) signum curved, with both ends bearing sparse and long spines.
Remarks. Based on the examined material, this species only occurs in Sri Lanka, from where the type was described (Walker 1859). However, because of the misidentifications as Ecpyrrhorrhoe machoeralis of E. damastesalis, records of the former are widely present in the literature (Wu et al. 1977, Intachat 1998, Lin et al. 2018, Wang 1980. The reported biology of E. machoeralis needs to be confirmed. The damaged female type material of E. machoeralis and lacks the original wing pattern because of its lack of scales, and the genital dissection is of low quality, which makes the identification and diagnosis of E. machoeralis difficult. Diagnosis. This species differs from all other species of the genus by the yellow forewing bearing rosy markings and mottled covering of rosy scales, the rosy markings on tornal area of the hindwing (Fig. 13), in the male genitalia (Fig. 29) by the longer and more strongly sclerotized arms on the juxta, with two groups of short and pointed spines on the anellus, in the female genitalia (Fig. 44) by the smaller and less developed antrum, and the smaller corpus bursae. Description (Fig. 13). Head. Frons and vertex yellow, frons with lateral white bands. Labial palpus yellow or orange, contrastingly white at base ventrally. Maxillary palpus yellow, pale terminally. Frons, vertex, labial and maxillary palpi sometimes mixed with rosy scales. Thorax. Dorsal side and tegula yellow, mixed with rosy scales; ventral side white. Wings. Forewing length: 10.0-12.0 mm. Forewing with termen weakly oblique; ground color yellow, with mottled covering of rosy scales forming indistinct markings except the totally rosy costa; antemedial line blurred, from 1/5 of costa oblique to 1/3 of 1A, then dentate to 2/5 of posterior margin; orbicular stigma nearly square; reniform stigma comma-shaped, thick and concave; postmedial line blurred, arched from 3/4 of costa to base of CuA 2 , connected with oval-shaped patch posterolateral of cell, and finally dentated to 2/3 of posterior margin; subterminal band with anterior part broad, inner margin serrated; fringe with basal half dark rosy and distal half pale yellow. Hindwing pale yellow; postmedial and subterminal lines rosy and serrated, but only with posterior part distinct; terminal line sometimes rosy; fringe as in forewing, with apex and tornus pale yellow. Underside: costal and terminal areas grey; reniform stigma and patch posterolateral of cell grey and distinct; postmedial line grey and faint. Abdomen. Yellowish to yellowish brown dorsally, the first two segments whitish, posterior margin of each segment paler. Dirty white ventrally.

Ecpyrrhorrhoe rosisquama
Male genitalia (Fig. 29). Uncus with basal 1/4 nearly triangular and naked. Valva slightly curved, of almost even width, apex obtusely rounded; sella thick and setose, thumb-shaped, bearing short spines on distal margin, extending ventrad; sacculus with distal 3/4 inflated into a broad and round protrusion. Juxta with base wide, distal 4/5 bifid into slender, long, and sclerotized arms; anellus with two groups of short and pointed spines (connected with distal arms of juxta in Fig. 29). Saccus rounded triangular. Phallus rather stout and straight, with interlaced spicules on vesical and cornuti present as a sclerite with dense and long spines.
Female genitalia (Fig. 44). Anterior apophysis ~ 2× the length of posterior apophysis. Antrum cup-shaped, weakly sclerotized, slightly wrinkled medially; colliculum moderately long and broad; ductus seminalis inserting to anterior end of colliculum and with a wide sclerite and a nearly semi-circular, strong sclerotized sclerite located opposite colliculum; ductus bursae ~ 3-4× diameter of corpus bursae, basal half bearing a slim sclerite. Corpus bursae globular; rhombic signum with maximal length < 1/2 of diameter of corpus bursae; second (posterior) signum composed of a pair of narrow and pointed sclerites without spines. Diagnosis. This species is similar to E. obliquata but can be best distinguished from it by the smaller size (forewing length: 7.0-9.0 mm), the pale brown reniform stigma and the indistinct and short streak of the forewing (Fig. 14), in the male genitalia (Fig. 30) by the excurved, finger-shaped and setose sella, the strongly sclerotized arms of the juxta, and anellus with a long spine and one or two short spines and in the female genitalia (Fig. 45) by the mostly tubular antrum.
Male genitalia (Fig. 30). Uncus with basal 2/5 nearly triangular and naked. Valva narrowly oval, costa straight or slightly concave and ventral margin curved, with apex rounded; sella excurved, thumb-shaped, thick, and setose, bearing thick setae on distal half; sacculus with distal 3/5 inflated into a broad and nearly triangular protrusion. Juxta with base wide, distal 3/4 bifid into slender, curved, sclerotized, and widely separated arms; anellus with a long spine and one or two short spines (attached to distal part of juxta in Fig. 30). Saccus rounded triangular. Phallus long and tapering, distal end spinulose and weakly sclerotized, with interlaced spicules cluster on vesica.
Distribution. China (Guangxi, Hainan, Jiangxi, Tibet, Yunnan). Etymology. The specific name is derived from the Latin exigu-(= short) and stria (= streak), referring to the short streak on forewings. Zhang, Li & Wang, 2004 Figs 15, 31, 46 Ecpyrrhorrhoe digitaliformis Zhang, Li & Wang, 2004: 318. Diagnosis. Forewing length: 9.0-14.0 mm. In appearance, Ecpyrrhorrhoe digitaliformis is indistinguishable from E. celatalis (Walker, 1859), but it can be distinguished from it in the male genitalia (Fig. 31) by the tapering and curved valva, the thumb-shaped, excurved and setose sella inflated distally, by the stout, sclerotized, finger-shaped dorsal protrusion of the sacculus, by the anellus with a string of minute spines and a coneshaped group of large spines (attached to distal phallus in Fig. 31), the modified distal ends of the arms of the juxta, and the shape of cornuti; in the female genitalia (Fig. 46), by the antrum with a vertical wrinkled area in the middle.

Ecpyrrhorrhoe digitaliformis
This species is closely related to E. brevis based on molecular data, and similar in appearance and male genitalia, but can be differentiated by the slender and excurved sella, the thick protrusion of sacculus, and the slender arms of juxta bearing a toothshaped process, as well as the characters mentioned above.
Male genitalia (Fig. 32). Uncus with basal half nearly triangular and naked. Valva curved and slowly tapering to rounded apex; sella thickly sclerotized, thumb-shaped, and densely setose; sacculus with distal 3/5 inflated into a broad protrusion bearing a slender finger-shaped process medially. Juxta with base wide, distal 4/5 bifid into stout and tapering, slightly curved and closely separated arms; anellus with a series of long spines standing on a long and curved base (attached to distal end of phallus in Fig.  32). Saccus broadly triangular. Phallus long and moderately stout, cornuti presented as a lancet-shaped sclerite connected with a hook-shaped, strong spine on apical end.
Distribution. China (Guangdong, Guangxi). Etymology. The specific name is derived from the Latin brevis (= short), referring to the short arms of juxta in the male genitalia.

Ecpyrrhorrhoe puralis (South, 1901) Figs 17, 33, 48
Pionea puralis South, 1901: 493. Diagnosis. Forewing length: 11.0-14.0 mm. Ecpyrrhorrhoe puralis is almost indistinguishable from E. longispinalis and E. biaculeiformis in appearance, but can be distinguished in the male genitalia (Fig. 33) by the small and excurved sella, much larger juxta with distal 2/3 bifid, anellus with comb-shaped spines (attached to distal phallus end of in Fig. 33), and in the female genitalia (Fig. 48)  Diagnosis. Forewing length: 11.0-14.0 mm. In appearance, Ecpyrrhorrhoe rubellalis resembles E. minnehaha, but can still be recognized by its larger forewing length, yellowish brown ground color of wings with yellow fringe, and more oblique antemedial line of forewing (Fig. 18); in the male genitalia (Fig. 34) by the broader valva with truncate tip, and the nearly spine-shaped, thin and short sella, by the phallus apically with a densely dentated, triangular projection and a thick spine bearing a broad and long, spinulose base on the anellus; in the female genitalia (Fig. 49) (Fig. 35) by the valva gradually broadening to the subapex, a hook-shaped sella, a small, sclerotized, ball-shaped sclerite bearing two small spines on opposite sides on the anellus; in the female genitalia (Fig. 50) by the antrum without sclerotized processes or triangular, wrinkled sclerites. Description (Fig. 19). Head. Frons pale yellow, with white lateral bands. Vertex pale yellow. Labial palpus dark yellow, contrastingly white at base ventrally. Maxillary palpus dark yellow, pale terminally. Thorax. Dorsal side dark yellow or yellowish brown, ventral side white. Legs white to pale yellow. Wings. Forewing length: 9.0-13.0 mm. Forewing bright yellow, termen moderately arched; antemedial line fulvous, outwardly curved from 1/4 of costa to 1/3 of posterior margin; orbicular stigma dot-shaped, small, sometimes indistinct; reniform stigma comma-shaped, concave; postmedial line from anterior 3/4 distinctively curved to middle of CuA 1 , then bending to 1/3 of CuA 2 , and finally undulated to 2/3 of posterior margin; terminal line and fringe bright yellow. Hindwing yellow, costal area white, postmedial line fulvous, slightly dentate curved, outward from 2/3 of M 1 to 1/2 of CuA 1 , arc-shaped, then bending inward along CuA 1 , reaching discocellular, then undulated to 2/3 of posterior margin; terminal line and fringe as in forewing. Abdomen. Pale yellow dorsally, black on distal part, white ventrally.
Male genitalia (Fig. 35). Uncus relatively thick, with basal half nearly triangular and naked. Valva curved and slowly broadening to rounded apex, with maximal width at sub-apex; sella hook-shaped with basal half densely setose; sacculus with distal 3/5 inflated into a triangular, rounded protrusion. Juxta with basal margin concave, distal half bifid into stout and pointed arms; anellus bearing a small and sclerotized ball, with two small spines on opposite sides (attached to distal end of phallus in Fig. 35). Saccus rounded triangular. Phallus long and slightly curved, cornuti presented as a long sclerite and a long and strong spine on apical end.
Female genitalia (Fig. 50). Anterior apophysis ~ 2× length of posterior apophysis. Lamella postvaginalis presented as a nearly trapezoidal sclerite. Antrum cup-shaped, strongly sclerotized, decorated with lots of small spines, those spines forming a circle, with a thumb-shaped, sclerotized process on the side of circle; colliculum narrow and moderately long; ductus seminalis connecting to anterior end of colliculum and with a short sclerite located opposite to colliculum; ductus bursae slender, length ~ 2× as long as diameter of corpus bursae, basal 1/3 bearing a slim sclerite. Corpus bursae globular; rhomboid signum with maximal length almost 1/3 as long as diameter of corpus bursae; second (posterior) signum nearly V-shaped bearing sparse and long spines. Distribution. China (Hubei, Hunan). Etymology. The specific name is derived from the combination of Latin long-and spinalis (= with spine), referring to the vesica with a long and thick spine. with many tiny spines on its basal 3/4 (attached to distal end of phallus in Fig. 36), in the female genitalia (Fig. 51) by the posterior part of the antrum looking like a pair of triangular sclerites. P. minnehaha (Pryer, 1877), P. rubicundalis Warren, 1896, P. rufipicta (Butler, 1880) and P. schenklingi Strand, 1918 are confirmed to belong in Ecpyrrhorrhoe. In the case of P. anpingialis Strand, 1918, the female genitalia of the holotype (♀, Anping, Formosa, IV.1912, H. Sauter Coll., Gen. präp. Gaedike NR: 9668 (SDEI)) does not have the diagnostic characters of Ecpyrrhorrhoe (absence of lamella antevaginalis, longitudinal stripe on ductus bursae, and second (posterior) signum) and is not congeneric with Ecpyrrhorrhoe, but its correct placement is unclear due to the lack of male material. The abdomens of the types of P. leucanalis Swinhoe, 1890 and P. suavalis (Walker, 1866) are lost. The genitalia slide of the type of P. fuscicostalis Swinhoe, 1894 is incorrect and may have been confused with that of Pyralidae Brit. Mus. Slide No. 8683, which is labelled with "incorrect abdomen? See 8683 for correct abdomen". The types of P. quadrigalis (Hering, 1901) and P. ignealis (Hampson, 1899) were not examined. Therefore, these six species are transferred to Ecpyrrhorrhoe temporarily, with their generic placement unconfirmed. Further study is needed to confirm their generic placement.
The genus Yezobotys Munroe & Mutuura, 1969 differs significantly in structure from Ecpyrrhorrhoe, and is more closely related to Anamalaia Munroe & Mutuura, 1969, based on examination the paratype material of Yezobotys ainualis Munroe & Mutuura, 1969 (Pyralidae Brit. Mus. Slide No. 19693 (NHMUK)). The generic characters of Yezobotys, the short and triangular uncus, the sacculus with finger-shaped process in male genitalia, and the strongly sclerotized lamella antevaginalis and postvaginalis in female genitalia, are extremely similar to those of Anamalaia Munroe & Mutuura. Thus, Yezobotys is restored as a valid genus.
According to the tree topology ( Fig. 1), the results of the phylogenetic analyses robustly support the monophyly of Ecpyrrhorrhoe in BI, but there is low support in ML (PP = 0.99, BS = 63) possibly caused by the missing data in the concatenated dataset. The genus Pagyda is the sister group of Ecpyrrhorrhoe (PP = 0.99, BS = 67), and Ecpyrrhorrhoe can be divided into three species groups (A clade, B clade and C clade), the B clade and C clade forming a sister group (PP = 1, BS = 49). The A clade (PP = 0.99, BS = 36), consisting of E. allochroa + E. damastesalis, can be distinguished from species in B clade and C clade by the following morphological characters: hindwing yellowish white without any lines or spot, instead of the brown postmedial line present on B clade and C clade; bifid arms of juxta short in male genitalia, ductus bursae without a slender, sclerotized, longitudinal sclerite in female genitalia. The B clade (PP = 0.95, BS = 29), consisting of seven species, can be differentiated by the transverse sclerite on the bottom of ductus seminalis in female genitalia. E. machoeralis, without molecular data and phylogenetic analysis, is assigned to B clade on the basis of morphological characters. The C clade (PP = 0.96, BS = 71), consisting of seven species, can be distinguished from species in A clade by a sclerotized and longitudinal stripe on ductus bursae in female genitalia, and distinguished from species in B clade by the absence of transverse sclerite on the base of ductus seminalis.
In this study, bootstrap values of the majority of the basal nodes are relatively low. Future research might utilize a broader sampling per species, fresher material more suitable for DNA studies, and additional genetic data to shed further light onto the phylogenetic relationships of this species complex.